THORPE, WILLIAM HOMAN

Thorpe played a major role in establishing ethology as a branch of academic biology in the English-speaking world after World War II. His ethological career was effectively his second, having made his reputation with diverse studies at Cambridge in entomology, on the strength of which he was elected a Fellow of the Royal Society in 1951. His discipline-building achievements on ethology’s behalf included the founding at Cambridge of an ornithological field station, where, alongside other research into the natural behavior of birds, he launched an enduring experimental project on the interplay of instinct and learning in song acquisition. Later decades saw him emerge as one of the leading voices against modern biology’s reductionist spirit, a theme he addressed in its scientific but also philosophical and, as a Quaker, religious dimensions. His several sides combined to political effect in his efforts to conserve natural sites and to safeguard animals from suffering.

From Entomology to Ethology. There was no tradition of farming in Thorpe’s family when he entered Jesus College, Cambridge, in 1921 to study agriculture. The son of an accountant, “Bill” (as he was always known) was raised in the southern English coastal towns of Hastings and Weston-super-Mare, where he came to share his parents’ pleasure in the outdoors and devotion to their Congregationalist faith. Like music, another lifelong passion dating from his childhood, natural history (especially birdwatching) became a pastime that suited a religiously minded and rather solitary boy. These tendencies combined with frequently poor health to make for an indifferent school career. Initially planning to make his way as a naturalist or farmer without the benefit of university training, he changed his mind on learning of the opportunities opening up in economic entomology. Taking advice, he decided on the Cambridge degree in agriculture as the best route, staying on for an entomologically focused Diploma year, then for a PhD (1929).

After a few years at the Farnham Royal Parasite Laboratory (part of the Imperial Institute of Entomology), Thorpe returned to Cambridge in 1932, taking up positions as a university lecturer in entomology and a tutor and Fellow of Jesus College. Already his main preoccupations as a research entomologist, engaged equally in theory, experiment, and field observation, had emerged. There was, first of all, an interest in the natural and artificial production of new biological “races,” and ultimately of new species, as different preferences for different kinds of food stabilized within a species. An example from Thorpe’s early research was the apparent differentiation of the ermine moth into biologically—though not (yet) structurally—distinct groups that fed and laid their eggs on different kinds of trees, apple or hawthorn. A second interest was in the role of parasites in controlling agriculturally destructive insects like ermine moths. A third interest was in figuring out the details of parasite respiration.

These related inquiries merged in the mid-1930s in a study of what Thorpe called “olfactory conditioning.” Having shown that a parasite had an inherited tendency to prefer a certain moth host, and that the parasite identified the larvae of this host by smell, Thorpe contaminated the larvae of a usually ignored kind of moth with the smell of the normal host to see whether he could induce the parasite to adopt the new host for egg laying. The experiment proved successful, with the parasite offspring, accustomed from birth to the smell of the new host, going on to have a weakened preference for the old host as egg-laying adults. For a time, Thorpe entertained the possibility that these host-preference studies might prove supportive of the “Lamarckian” doctrine of the inheritance of acquired characters. Although he eventually decided that they did not, animal learning—for decades the domain of the psychologists—came increasingly to dominate his research. Discovering the work of the Austrian zoologist Konrad Lorenz, who under the banner of “ethology” was refashioning learning, and with it instinct, as zoological topics, confirmed Thorpe in his new direction.

Establishing Ethology in Britain. After World War II, Thorpe committed himself to a new science, ethology, as well as a new religion, Quakerism. In neither case was there dramatic conversion. He had been attending meetings of the Society of Friends since his Farnham Royal days. When the war came, he registered, on the Quaker model, as a conscientious objector, though he made himself useful as an expert in eliminating the agricultural pests that threatened Britain’s ability to feed itself. His turn toward ethology, and toward the study of the instinctive and learned behaviors of birds—Lorenz’s favored organisms—was similarly an extension of gradual. For all that the immediate postwar period saw Thorpe energetically proselytizing for ethology, he was also immersed in his most innovative research thus far on the respiratory physiology of insects. By the early 1950s, however, the disciplinary change was complete: a remarkable, midlife transformation in someone who, as his new status as a Fellow of the Royal Society demonstrated, had already made his mark.

Thorpe had much to offer ethology, particularly with respect to its institutional development. A science with a small but expanding institutional life in continental Europe before the war, ethology, like war-torn Europe itself, needed reconstruction. Niko Tinbergen, the Dutch zoologist commonly regarded as ethology’s co-founder, was attempting this at the University of Leiden. Lorenz, however, who had been captured by the Russians and interred as a prisoner of war, had no institutional base in Austria when he finally returned there in 1948. Rumors— not without a certain basis—that Lorenz had had a Nazi past were not yet widespread. At least they do not appear to have been an issue for Thorpe as he sought to give ethology a new home in Britain. As with most complex undertakings, the motivations underpinning this one were mixed and mostly obscure. Whatever ambitions, scientific or spiritual, gave Thorpe his sense of mission, what is undoubted is that he admired Lorenz’s prewar papers deeply, especially those on the kind of instinctconstrained learning that Lorenz’s translators called “imprinting,” and which Thorpe had retrospectively recognized as at work in his moth parasites.

Orchestrating ethology’s return engaged him on several fronts. During the war he joined the Institute for the Study of Animal Behaviour, whose members and Bulletin sought to promote just the sort of naturalistic, objective studies the ethologists prized. After the war’s end, he began corresponding with Tinbergen, accepting Tinbergen’s invitation to be an editor of Tinbergen’s new journal for ethological research, Behaviour. When the news came in 1948 that Lorenz was alive and back home near Vienna, Thorpe made the journey, exploiting some personal connections to arrange some emergency research funding for Lorenz. Working again with Tinbergen, Thorpe began planning a conference that, as a meeting in Cambridge in 1949 and subsequently in its published proceedings, would do much to publicize ethological aims, methods, theories, and personalities. None of the personalities came larger than Lorenz's; and one of the meeting’s many successes was the reunion at Thorpe’s Cambridge home between Lorenz and Tinbergen.

Although not everyone in Cambridge zoology was as impressed with ethology as was Thorpe, he managed the following year to start an ornithological field station on four acres of woods and meadows recently acquired by the university in the nearby village of Madingley. There was talk for a time of hiring Lorenz as curator. The position ultimately went to Robert Hinde, who, fresh from doctoral studies with Tinbergen at Oxford, was soon at work setting up, stocking, and maintaining the aviaries that became the mainstay of the station’s research. Between them, the Lorenz-inspired Thorpe and the Tinbergeninspired Hinde trained generations of Cambridge graduate students in ethology. Acutely aware of differences between psychological and ethological approaches, the two men nevertheless reached out to psychological colleagues, in Thorpe’s case most notably in his extraordinary synthesis of studies in both traditions, Learning and Instinct in Animals(1956), and in a famous seminar series run with the Cambridge psychologist Oliver Zangwill.

Experiments in Song Learning. Like his embrace of ethology more generally, Thorpe’s turn toward bird behavior, and in particular song acquisition in the chaffinch, had its roots in his pre-ethological work. As a student in the 1920s he had been an avid bird-watcher, cofounding the Cambridge Bird Club and even publishing ornithological papers. Chaffinches were and are plentiful in Cambridge, indeed throughout Britain. What most likely drew them to Thorpe’s attention in the 1930s, however, was well-known research suggesting that chaffinches exhibited behavior patterns that—as in the moths he was then studying—could be transmitted stably from one generation to the next without being entirely genetically fixed. The behavior patterns in question were song versions or “dialects,” whose maintenance in separate areas of a region of Russia was thought to arise from chaffinches having learned the dialects sung around them as juveniles and then returning to their birthplaces for mating. Here instinct and experience seemed to mix in just the ways that Lorenz had made central to ethologists’ concerns.

After the war, as Thorpe developed his plans for an experimental study of song learning in the chaffinch, he became aware of a new technology apparently custommade for his purposes. Fitting onto a desktop, the sound spectrograph, or Sonagraph to use its commercial name, produced visual displays showing the distribution of frequencies and intensities making up a complex sound. Although invented at Bell Laboratories as a secret wartime project (enabling, for instance, the identification at a distance of different vessels by their underwater acoustics), the device was publicized at war’s end as a boon for the deaf, helping them to read incoming telephone calls and reply with speech improved through spectrograph-aided education. Almost incidentally, the paper introducing the spectrograph included bird-song spectrograms, made to test how well it handled rapid changes of pitch. Among the ornithologists who saw the potential was Thorpe, who, in the early 1950s, made use of a British military spectrograph before acquiring his own.

His 1954 paper in Nature, “The Process of Song-Learning in the Chaffinch as Studied by Means of the Sound Spectrograph,” gave notice to the scientific community that Cambridge was now home to a novel research program on the interaction of learning and instinct. The basic experimental method was straightforward. With spectrographic snapshots of the songs of normally raised local chaffinches serving to define “normal” song, Thorpe and the station’s first graduate student, Peter Marler, systematically varied the conditions under which birds were raised in order to see—literally, thanks to the spectrograph—how the adult songs of these birds came to differ from normal song, and so how sensitive the song acquisition process was to changes in experience. To take the simplest example, individuals reared in acoustic isolation almost from birth were found to sing crude but still chaffinchlike songs. These experiments appeared to reveal the species’ innate song, before experience elaborated and refined it.

Courtesy of the engineering prowess of Bell Labs and its military and commercial agenda, a whole new class of animal behaviors—vocalizations—was now open to quantitatively robust ethological investigation. Between them, Thorpe and Marler exploited the new opportunities to the fullest. Before leaving Madingley in 1957 for the University of California at Berkeley, where he went on to develop the song-learning paradigm using a local species, the white-crowned sparrow, Marler published classic papers on, for instance, the relationship between the biological function of bird calls and their spectrographically revealed acoustic structures (ethological doctrine of the day had it that signal structures were arbitrary). Thorpe summarized Marler’s conclusions and much else in a compact 1961 book, Bird-Song: The Biology of Vocal Communication and Expression in Birds. It would be his last major contribution to a research lineage that, first under Marler at Berkeley and the Rockefeller University, then under Marler’s former students at Rockefeller and elsewhere, continues to yield textbook-altering research.

Antireductionism. Animal behavior is no more intrinsically “reductionist” or “holist” than any other area of biology. It is nevertheless true that, from the later nineteenth century, an emphasis on the (especially purposeful) activity of animals and its evolutionary importance has often appealed to those wishing, for one reason or another, to strike a blow against the idea that science has revealed life to be mechanism all the way down and all the way up. The spiritually questing Thorpe belongs firmly in this tradition. His early work on biological races and the emergence of new feeding habits grew out of sympathetic interest in one of the great antimechanical themes of modern biology, the inheritance of acquired characters. Among animal psychologists, where learning through trial and error defined the mechanical extreme, dissenters tend to concentrate on problems whose solutions demand that animals perceive and act on relationships. Alongside his song research, such “insight” learning in birds became one of Thorpe’s major ethological preoccupations.

After 1961, as Thorpe entered his sixties, he continued with his experimental research. There was, however, a noticeable shifting in ambition, as he turned increasingly from the business of doing animal behavior biology, for an audience of scientific peers, to the business of reflecting on it for the benefit of the general public. Two lecture-series-turned-books, Biology and the Nature of Man(1962) and Science, Man, and Morals(1965), brought into the open the largely implicit antireductionist thrust of his previous work, as well as the all but invisible Christian imperatives to which it answered. Exhibiting humankind as the product of an evolutionary process regularly transcending mere mechanism, Thorpe engaged his considerable synthetic talents to ranging over immense tracts of biology, psychology, philosophy, and theology. With such performances, it is little wonder that when the man of letters Arthur Koestler organized a soon-famous 1968 symposium on the life sciences “beyond reductionism,” he secured Thorpe’s participation.

The culmination of Thorpe’s efforts along these lines was Animal Nature and Human Nature(1974), a long tome born of his lectures in the distinguished natural-theological Gifford series at St. Andrews between 1969— the year of his formal retirement from the Chair of Ethology created for him three years previously—and 1971. Old-fashioned in spirit but up to date in its materials, the book built toward its uplifting vision of evolutionary progress by way of everything from the difference between life and nonlife to man as a religious animal. There was, however, a new challenge to be faced: a popular reductionist tract on biology, Chance and Necessity (1971), by the French molecular geneticist Jacques Monod. No less unsettling from Thorpe’s perspective was the emergence of a new “sociobiology,” bent not just on reducing animal social behavior to genetic mechanisms but on bypassing ethology in doing so. Responding to these developments led Thorpe to write his final books, respectively, Purpose in a World of Chance(1978) and the valedictory The Origins and Rise of Ethology(1979). He died peacefully in a nursing home in 1986.

Legacies. What, aside from the song-learning research tradition, endures of Thorpe’s achievements? His articles and books are not much read nowadays; and ethology as an identifiable discipline did not long survive in the sociobiological era. But the ethological attitude remains a live option, not least because several of the institutions that Thorpe helped establish still flourish, notably the Madingley station, long since grown into a subdepartment of animal behavior at Cambridge. Perhaps the legacy touching the most people, however, lies with what Thorpe once, in the Quaker journal Friend(he was a frequent contributor), called “Man’s Responsibilities To and For Nature.” In his view, evolution’s gifts to humankind brought with them a duty of care. No mere thinker on this matter, he campaigned energetically for decades on behalf of the conservation of wildlife areas and—with rather more delayed success—animal welfare in laboratories and factory farms. Rarely have science and religion reconciled so beneficently.

BIBLIOGRAPHY

The papers of William Homan Thorpe are maintained at the, Department of Manuscripts and University Archives, Cambridge University Library, Cambridge, U.K . A very full bibliography of Thorpe’s writings can be found in Robert Hinde’s obituary notice (see below).

“Types of Learning in Insects and Other Arthropods.” Parts 1, 2, and 3. British Journal of Psychology 33 (1942–1943): 220–235; 34 (1943–1944): 20–31, 66–76. The first of several synthetic surveys of animal learning.

OTHER SOURCES

Boakes, Robert. From Darwin to Behaviourism: Psychology and the Minds of Animals. Cambridge, U.K.: Cambridge University Press, 1984. An excellent introduction to animal psychology as Thorpe came to know it, including insight learning.

Burkhardt, Richard W., Jr. Patterns of Behaviour: Konrad Lorenz, Niko Tinbergen, and the Founding of Ethology. Chicago: University of Chicago Press, 2005. An outstanding history of ethology, with much of interest on Thorpe, some of it based on interviews.

Durant, John R. “The Making of Ethology: The Association for the Study of Animal Behaviour, 1936–1986.” Animal Behaviour 34 (1986): 1601–1616. An invaluable study of this organization and Thorpe’s role in particular.

Gillispie, Neal C. “The Interface of Natural Theology and Science in the Ethology of W. H. Thorpe.” Journal of the History of Biology 23 (1990): 1–38. The most extensive discussion of this topic currently available.

Hinde, Robert A. “William Homan Thorpe, 1902–1986.” Biographical Memoirs of Fellows of the Royal Society 33 (1987): 620–639. The fullest of the obituary notices, with excellent summaries of Thorpe’s scientific writings.

Marler, Peter. “Science and Birdsong: The Good Old Days.” In Nature’s Music: The Science of Birdsong, edited by Peter Marler and Hans Slabbekorn. Boston: Elsevier, 2004. A concise and authoritative overview of the research program Thorpe began; includes a CD with Thorpe’s own chaffinch recordings.

Wilson, David A. H. “Animal Psychology and Ethology in Britain and the Emergence of Professional Concern for the Concept of Ethical Cost.” Studies in History and Philosophy of Biological and Biomedical Sciences 33 (2002): 235–261. A pioneering article, placing Thorpe’s animal welfare activities in context.

Gregory Radick

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